“Like a fish out of water: How does the African lungfish

survive the changing seasons?”

Emily Magkourilou
School of Biological Sciences, University of Aberdeen, Scotland

There are four extant species of African lungfish; all belonging to the genus Protopterus [1].
Phylogenomic studies have shown that lungfishes are the closest living relatives to
tetrapods, making them a model taxon to study the evolutionary transition from water to
land [1][2]. African lungfishes in particular are also a model for morpho-physiological
adaptations to environmental challenges, as they manage to survive in areas of intertropical
Africa exposed to low oxygen levels, high temperatures, and most importantly seasonal
drought [3]. In order to avoid hypoxia and desiccation when the water basins dry up, African
lungfishes use their elongated bodies and threadlike appendages to burrow and form a
cocoon around their body in a process known as estivation [2][3]. In this 3-phased dormant
state, the animals can survive without food and water from a few months up to five years
[2][3]. During the induction phase (first 6-8 days), cocoon formation takes place by
hyperventilation and mucus secretion [3]. Also, osmoregulation is enhanced as mRNA
expression of growth hormone and prolactin increases [2]. Next, the maintenance phase is
characterised by a metabolic depression of more than 50% in non-vital organs [4]. This is
achieved via a coordinated suppression of the rate of cellular ATP consumption [5].
Meanwhile, muscle tail digestion and increased lipid metabolism enable survival on internal
fuel reserves [4]. There is also a readjustment of the metabolic pathway from ammonotelism
to the less toxic ureotelism [6]. Down-regulation of respiratory and cardiovascular rates also
takes place [3]. All African lungfish species are obligate air breathers, with two lungs
resulting from a modified swim bladder, and reduced but functioning gills [3]. During
maintenance, the rate of oxygen consumption decreases whilst the mucus covered gills
become redundant [3]. On the other hand, the lungs expand and vascularise allowing the
animal to depend entirely on pulmonary breathing [3]. At the end of maintenance, there is
an up-regulation of transcription, translation, and glycolytic capacities in preparation for the
last phase, arousal [7]. Upon contact with water, the animal enters arousal leaving the
cocoon and slowly recovering muscle activity in order to forage for tissue repair and growth
[2]. After arousal, accumulated urea is excreted by transcriptional activation of putative urea
transporters (lfUT) found in the skin [8]. Overall, African lungfishes have evolved a wide
range of morphological, respiratory, and metabolic adaptations. These are tightly
coordinated and allow the animals to survive the extremes of the African dry and wet
tropical climate.
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